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CLASSIFICATION & EVOLUTION Humans classify almost everything, including each other. This habit can be quite useful. For example, when talking about a car someone might describe it as a 4-door sedan with a fuel injected V-8 engine. A knowledgeable listener who has not seen the car will still have a good idea of what it is like because of certain characteristics it shares with other familiar cars. Humans have been classifying plants & animals for a lot longer than they have been classifying cars, but the principle is much the same. In fact, one of the central problems in biology is the classification of organisms on the basis of shared characteristics (i.e., simplesiomorphies vs. synapomorphies). As an example, biologists classify all organisms with a backbone as "vertebrates." In this case the backbone is a characteristic that defines the group. If, in addition to a backbone, an organism has gills & fins it is a fish, a subcategory of the vertebrates. This fish can be further assigned to smaller & smaller categories down to the level of the species. The classification of organisms in this way aids the biologist by bringing order to what would otherwise be a bewildering diversity of species. (There are probably several million species - of which about one million have been named and classified.) The field devoted to the classification of organisms is called Taxonomy [Gk. taxis, arrange, put in order + nomos, law]. The modern taxonomic system was devised by Carolus Linnaeus (1707-1778). It is a hierarchical system since organisms are grouped into ever more inclusive categories from species up to kingdom. Figure 1 illustrates how four species are classified using this taxonomic system. (Note that it is standard practice to underline or italicize the genus & species names.) KINGDOM PHYLUM CLASS ORDER FAMILY GENUS SPECIES Animalia Chordata Mammalia Primate Carnivora Hominidae Canidae Homo Canis sapiens lupus (human) (wolf) Arthropoda Insecta Hymenoptera Apidae Apis mellifera (honeybee) Plantae Angiospermophyta Monocotyledoneae Liliales Liliaceae Alium sativum (garlic) Figure 1 In the 18th century most scientists believed that the Earth and all the organisms on it had been created suddenly in their present form as recently as 4004 BC. According to this view, Linnaeus' system of classification was simply a useful means of cataloging the diversity of life. Some scientists went further, suggesting that taxonomy provided insight into the Creator's mind ("Natural Theology"). This view of taxonomy changed dramatically when Charles Darwin published On The Origin of Species in 1859. In his book Darwin presented convincing evidence that life had evolved through the process of Natural Selection. The evidence gathered by Darwin, & thousands of other biologist since then, indicates that all organisms are descended from a common ancestor. In the almost unimaginable span of time since the first organisms arose (about 3.5 billion years) life has gradually diversified into the myriad forms we see today. As a consequence of Darwin's work it is now recognized that taxonomic classifications are actually reflections of evolutionary history. For example, Linnaeus put humans & wolves in the class Mammalia within the phylum Chordata because they share certain characteristics (e.g. backbone, hair, homeothermy, etc.). We now know that this similarity is not a coincidence; both species inherited these traits from the same Common Ancestor. In general, the greater the resemblance between two species, the more recently they diverged from a common ancestor. Thus when we say that the human & wolf are more closely related to each other than either is to the honeybee we mean that they share a common ancestor that is not shared with the honeybee. Wolf Figure 2 The vertical axis in this figure represents time. The point at which two lines separate indicates when a particular lineage split. For example, we see that mammals diverged from reptiles about 150 million years ago. The most recent common ancestor shared by mammals & reptiles is indicated by the point labeled A. The horizontal axis represents, in a general way, the amount of divergence that has occurred between different groups; the greater the distance, the more different their appearance. Note that because they share a fairly recent ancestor, species within the same taxonomic group (e.g. the class Mammalia) tend to be closer to each other at the top of the tree than they are to members of other groups. Several types of evidence can elucidate the evolutionary relationship between organisms, whether in the form of a taxonomic classification (Fig. 1) or a tree (Fig. 2). One approach, as already discussed, is to compare living species. The greater the differences between them, the longer ago they presumably diverged. There are, however, pitfalls with this approach. For example, some species resemble each other because they independently evolved similar structures in response to similar environments or ways of life, not because they share a recent common ancestor. This is called Convergent Evolution because distantly related species seem to converge in appearance (become more similar). Examples of convergent evolution include the wings of bats, birds & insects, or the streamlined shape of whales & fish. At first glance it might appear that whales are a type of fish. Upon further examination it becomes apparent that this resemblance is superficial, resulting from the fact that whales & fish have adapted to the same environment. The presence of hair, the ability to lactate & homeothermy clearly demonstrate that whales are mammals. Thus, the taxonomist must take into account a whole suite of characteristics, not just a single one. The fossil record can also be helpful for constructing phylogenetic trees. For example, bears were once thought to be a distinct group within the order Carnivora. Recently discovered fossils, however, show that they actually diverged from the Canidae (wolves, etc.) fairly recently. The use of fossils is not without its problems, however. The most notable of these is that the fossil record is incomplete. This is more of a problem for some organisms than others. For example, organisms with shells or bony skeletons are more likely to be preserved than those without hard body parts. The Classification & Evolution of Artificial Organisms In this exercise you will develop a taxonomic classification & phylogenetic tree for a group of imaginary organisms. On the last page of this handout there are pictures of the 14 "living" & 58 "fossil" species that you will use. Take a look at the pictures & note the variety of appendages, shell shapes, patterns, etc. Each species is identified by a number rather than a name. For fossil taxa there is also a number in parentheses indicating the geological age of each specimen in millions of years. Most of the fossils are extinct, but you will notice that a few are still living (e.g. species #24 is found among the living forms, but there is also a 2 million year old fossil of #24 in our collection). The purpose of this exercise is to illustrate the principles of classification & some of the processes of evolution (e.g. Convergent Evolution). We do these exercises with artificial organisms so that you will approach the task with no “preconceived” notion as to how they should be classified. This means that you will have to deal with problems such as convergent evolution just as a taxonomist would. With real organisms you would probably already have a pretty good idea of how they should be classified & thus miss some of the benefit of the exercise. Exercise 1: The Taxonomic Classification of Living Taxa Carefully examine the fourteen living species & note the many similarities & differences between them. Create a hierarchical classification of these species, using the format in Figure 3. Instead of using letters (A, B, ...), as in this example, use the number of each species. Keep in mind that Figure 3 is just a hypothetical example. Your classification may look quite different than this one. GENUS 1 A G PHYLUM CLASS 1 ORDER 1 FAMILY 1 FAMILY 2 GENUS 2 GENUS 3 GENUS 4 H D B J L CLASS 2 ORDER 2 ORDER 3 FAMILY 3 FAMILY 3 GENUS 5 GENUS 6 E K C F I Figure 3 The first step in this exercise is to decide which species belong in the same genus. Species within the same genus share characteristics not found in any other genera. The taxa numbered 19 & 20 are a good example; they are clearly more similar to each other than either is to any of the other living species so we would put them together in their own genus. Use the same procedure to combine the genera into families. Again, the different genera within a family should be more similar to each other than they are to genera in other families. Families can then be combined into orders, orders into classes & so on. Exercise 2. The Comparative Approach to Phylogenetic Analysis Construct a tree based only on your examination of the 14 living species. This tree should reflect your taxonomic classification. For example, let us say you have put species A & G into the same genus because you think they evolved from a common ancestor (x). Their part of the tree would look like the diagram on the right. A When there are three or more species in a genus you must decide which two of the E species share a common ancestor not shared by the other(s). This diagram indicates that species E & K are more closely related to each other than either is to C. We y hypothesize that E & K have a common ancestor (y) that is not shared by C. Similarly, two genera that more closely resemble each other than they do other genera presumably share a common ancestor. Thus, even in the absence of a fossil record it is possible to develop a tree. We can even infer what a common ancestor like y might have looked like. G x K C z Exercise 3. The Tree Construct a tree for these taxa. Beginning with the oldest fossils, arrange the taxa according to their evolutionary relationship. For your reconstruction, use the living taxa (remember some may be contained in the fossil record) as well as taxa 15, 33, 36, 43, 44, 50, 52, 54, 57, 60, 73, 74, & 77. *For extra credit, try reconstructing a tree using the entire data set. Hints, Suggestions, & Warnings a. Draw lines to indicate the path of evolution. Start with the oldest fossil & work your way to the present i.e., use a timescale as your “Y” axis. b. Branching should involve only two lines at a time: Like this YES Not this NO c. Some living forms are also found in the fossil record! d. There are gaps in the fossil record for some lineages. Also, some species went extinct without leaving any descendants (this happens quite often in “real life”). e. The taxa were numbered at random; the numbers provide no clues to their evolutionary relationships. f. There is only one correct phylogenetic tree in this exercise. Start with the most primitive form (#73) & gradually modified it using a process that mimics evolution in real organisms. Group Questions 1. Provide answers to Exercises 1 & 2. 2. Provide a diagram describing the relationship among your living & fossil Taxa. 3. Provide a justification as to why you constructed the tree the way you did i.e., are all your groups, Genera & Families, Monophyletic? What characters were important in your analysis? 4. What sorts of problems were encountered i.e., are there any regions of your tree that need additional fossil data to reconstruct the relationships? 5. Did any of the taxa go extinct? How do they contribute to your understanding of the Evolution of these species? 6. Are there any species, or groups, that show Convergent Evolution? Did these hinder your efforts to contruct a tree, why or why not? 7. Would you describe the process of classification as difficult or easy, explain your answer? How much more difficult would your job have been if you had been working from the present to the past? LIVING Taxa FOSSIL Taxa (numbers in parentheses indicate age in millions of years)